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PubMed2002

Egg of the Karner Blue butterfly (Lycaeides melissa samuelis): morphology and elemental analysis.

Most insect eggshells are ornately sculptured; that of the Karner Blue butterfly, Lycaeides melissa samuelis, exhibits a series of interwoven ridges and depressions. Scanning electron microscopic views of the shell show that the patterning resides in the outer chorion, while the inner vitelline membrane is relatively flat and featureless. We here describe the morphology of the egg and introduce a physical technique, use of a Dynamitron accelerator, to identify and localize elements in the eggshell. Most elements present are represented in the chorion, but sulfur appears restricted to the vitelline membrane. The micropyle is particularly rich in calcium and, in unhatched eggs, phosphorus as well.

Animals↗
PubMed1995

Eyespot development on butterfly wings: the epidermal response to damage.

Eyespot colour patterns decorate the wings of many butterfly species. The eyespot is specified in the early pupal epidermis by signals from a central "focus," and it has been suggested that the focus is the source of a diffusible morphogen gradient. We show that ectopic eyespots can be induced in nonfocal positions throughout the distal, but not the proximal, wing epidermis of Bicyclus anynana by mild epidermal damage inflicted at 12-18 hr (into a 6- to 7-day pupal period). Damage may lower, locally and transiently, the threshold for response to morphogen. Here, we have tested two predictions of the gradient model. As predicted, mild damage close to the focus (parafocal) locally extends the eyespot, making it markedly asymmetrical, even after early operations (1 or 6 hr), when remote epidermis is non-responsive. Early parafocal operations also have an unexpected result, however, reducing the extent of the eyespot in other directions, perhaps through a long-range "wound effect" on the signaling activity of the focal cells. The model also correctly predicts that increasing the severity of a nonfocal operation will prolong the transient damage and hence give ectopic eyespots after early (1- or 6-hr) operations. We do not, however, find the expected size increase in ectopics induced by the later (12 or 18 hr) severe damage. Similarly, we demonstrate the predicted effect of early (1 or 6 hr) mild damage in increasing the response to a subsequent (18 hr) operation at the same position. The gradient model is therefore supported by most aspects of eyespot induction in response to epidermal damage.

Animals↗
PubMed1999

Molecular phylogeny of swallowtail butterflies of the tribe Papilionini (Papilionidae, Lepidoptera).

Swallowtail butterflies of the tribe Papilionini number about 225 species and are currently used as model organisms in several research areas, including genetics, chemical ecology and phylogenetics of host plant utilization and mimicry, mechanisms of speciation, and conservation. We have inferred phylogenetic relationships for a sample of 18 species of the genus Papilio (sensu lato) and five outgroup taxa by sequencing two stretches of mitochondrial DNA that correspond to segments 12886-13370 and 12083-12545 of Drosophila melanogaster mitochondrial DNA and consist of sections of the genes for the large ribosomal RNA and subunit 1 of NADH-dehydrogenase. Our data support the monophyly of Papilio and, within it, of several traditionally recognized subgroups. Species belonging to groups that utilize primarily Rutaceae as larval foodplants form two clusters, corresponding to Old World and American taxa, respectively, while two previously recognized clades-of American and South Asian-Austronesian origin-whose members were known to feed mostly on Lauraceae and Magnoliaceae, are observed to form a clade. The sister group of Papilio is found to be the South Asian genus Meandrusa, which also happens to feed on Lauraceae. The latter plant family is therefore the probable larval host of the ancestor Papilio and the shift to Rutaceae (which four-fifths of extant Papilio species use as foodplants) is more likely to have occurred only after the initial diversification of the genus.

Animals↗
PubMed1981

Fine structure of the stretch receptor in the bursa copulatrix of the butterfly, Pieris rapae crucivora.

A pair of multipolar stretch-receptive neurons were found in the bursa copulatrix of the female cabbage white butterfly, Pieris rapae crucivora. The cell body of each neuron, about 10 micrometers in diameter, lies on the edge of the muscular region in the antero-lateral wall of the corpus bursae. No special accessory structure, and as a receptor muscle, is associated with the neuron. The several dendrites extend radially into the muscle layer. The dendrites are ensheathed except for their terminal tips, and, on their course, they anchor repeatedly on the epithelial cells or the muscle fibers in such a manner that their basement membranes fuse together. While the ensheathed dendrite is usually 0.1-0.2 micrometer in diameter, it often forms 1-2 micrometers varicosities especially at anchor sites, so that it looks like a varicose, or beaded, chain. The varicosities contain a number of mitochondria, but only microtubules are found in the fine interconnecting parts of the dendrite. The naked dendritic tips terminate in the basement membrane of the epithelial cell. The varicosities, as well as naked tips, seem to be important for stimulus transduction in the sensory cell of this type.

Animals↗
PubMed1976

Polymorphism and predictability at the alpha-glycerophosphate dehydrogenase locus in Colias butterflies: gradients in allele frequency within single populations.

Heterozygosity at the alpha-glycerophosphate dehydrogenase locus of five species of Colias butterflies is widespread in montane populations; alpine and lowland populations are not heterozygous. Within a single demographically characterized population of C. meadii where the population extends from alpine down into montane habitats, a marked cline in allele frequency is seen. Such within-population clines suggest the involvement of strong selection across the marked ecological interface. Thermal factors are the most likely causative agents, but associative overdominance is not excluded.

Acclimatization↗
PubMed1984

Termination profiles of photoreceptor cells in the larval eye of the swallowtail butterfly.

Profiles of the axonal endings of seven retinular cells (R1-7) from each of six stemmata (I-VI) of the butterfly Papilio xuthus were examined. Their axons enter the brain anteroventrally and project dorsally along the brain's lateral surface before entering the optic neuropil. Larval optic neuropil has two distinct areas, a (distal) lamina and a (proximal) medulla, connected by a chiasma. Intracellular administration of cobalt shows that distal retinular cells (R1-3 of stemmata I-IV, R1-3 and R6 of stemmata V and VI) terminate as short axons with plug or toothbrush-like endings in the lamina. Proximal retinular cells (R4-7 of stemmata I-IV, R4, R5 and R7 of stemmata V and VI) terminate as long axons in the chiasma or in the medulla. The long axons have fine branches in the lamina, chiasma and medulla. The two proximal cells (R5 and R7) are distinguished by their deeper endings and longer branches. The terminals of photoreceptors correspond to their photoreceptive properties and provide further evidence for the functional specialization of the photoreceptors.

Animals↗
PubMed1993

The relationship between the Pgi locus and the ability to fly at low temperatures in the monarch butterfly Danaus plexippus.

Differences in genetic structure of samples of monarch butterflies caught at different times of day have been reported previously. This paper compares differences in allele and heterozygote frequencies at the Pgi locus between animals flying early and those flying late in outdoor flight cages and between animals able and animals unable to fly at a constant low temperature. There were consistent effects across a number of tests and in comparisons with field data, especially in males. Animals with the M allele were more likely to be able to fly at low temperatures, to become active early in outdoor flight cages, and to be caught early in the field. Also, differences were observed between males and females in the effect of allele on flight activity.

Alleles↗
PubMed2002

Feminization of genetic males by a symbiotic bacterium in a butterfly, Eurema hecabe (Lepidoptera: Pieridae).

Wolbachia are symbiotic bacteria found in many arthropods and filarian nematodes. They often manipulate the reproduction of host arthropods. In the present study, female-biased sex-ratio distortion in the butterfly Eurema hecabe was investigated. Breeding experiments showed that this distorted sex ratio is maternally inherited. When treated with tetracycline, adult females of the thelygenic line produced male progeny only. After PCR using Wolbachia-specific primers for the ftsZ gene a positive result was seen in the thelygenic females, but not in male progeny from tetracycline-treated females, or individuals from a Tokyo population with normal sex ratio and reproduction. Cytological observations showed that thelygenic females lack the sex chromatin body (W chromosome). The results strongly suggest that the sex-ratio distortion in E. hecabe is due to feminization of genetic males by Wolbachia.

Animals↗
PubMed2004

A butterfly's chemical key to various ant forts: intersection-odour or aggregate-odour multi-host mimicry?

Deception is a crucial yet incompletely understood strategy of social parasites. In central Europe, the Mountain Alcon Blue, Maculinea rebeli, a highly endangered butterfly, parasitises several Myrmica ant species. Caterpillars gain access to host nests probably by faking the ants' odour. We analysed gas chromatography-mass spectrometry data of body surface hydrocarbons of pre-adoption and hibernated larvae of Maculinea rebeli and of their host species Myrmica sabuleti and M. schencki. Data were ordinated by different methods, based on similarities in the relative quantities of compounds between chromatograms. The two Myrmica species exhibit species-specific profiles. The Maculinea rebeli pre-adoption larva has a complex profile that simultaneously contains species-specific substances of the two investigated host species. This evidence leads to the interpretation that, in central Europe, Maculinea rebeli is predisposed for multi-host use by the chemical signature of its pre-adoption larva. The Maculinea rebeli larva clearly does not rely on an "intersection-odour" of compounds common to all host ant species, but synthesises an "aggregate-odour" containing specific compounds of each of the investigated hosts. We term this previously unknown chemical strategy "aggregate-odour multi-host mimicry".

Animal Feed↗
PubMed2006

A male gift to its partner? Cyanogenic glycosides in the spermatophore of longwing butterflies (Heliconius).

Males of several insect species transfer nuptial gifts to females during mating, typically in the form of a protein-rich spermatophore. In chemically defended species, males could potentially enhance such a gift with chemicals that help protect the female, her eggs, or both. This was shown for lepidopteran species that accumulate pyrrolizidine alkaloids. Most Heliconius butterflies are presumably protected from predators by virtue of de novo synthesized and/or sequestered cyanogenic glycosides. Males of Heliconius species are known to transfer nutritional gifts to the females but whether defensive chemicals could also be transferred is not known. To ascertain whether transfer of cyanogens occurs, we dissected freshly mated females from nine different Heliconius species and analyzed spermatophores for cyanogenic glycosides. We found cyanogens in the spermatophores of all nine species. This is the first time cyanogenic glycosides are reported in the spermatophores of arthropods. We discuss the implications of these findings for Heliconius biology and for other cyanogenic insects as well. We suggest that chemically defended species commonly lace their nuptial gifts with defensive chemicals to improve gift quality.

Animals↗
PubMed2000

Six opsins from the butterfly Papilio glaucus: molecular phylogenetic evidence for paralogous origins of red-sensitive visual pigments in insects.

It has been hypothesized that the UV-, blue-, and green-sensitive visual pigments of insects were present in the common ancestor of crustaceans and insects, whereas red-sensitive visual pigments evolved later as a result of convergent evolution. This hypothesis is examined with respect to the placement of six opsins from the swallowtail butterfly Papilio glaucus (PglRh1-6) in relationship to 46 other insect, crustacean, and chelicerate opsin sequences. All basal relationships established with maximum parsimony analysis except two are present in the distance and maximum likelihood analyses. In all analyses, the six P. glaucus opsins fall into three well-supported clades, comprised, respectively, of ultraviolet (UV), blue, and long-wavelength (LW) pigments, which appear to predate the radiation of the insects. Lepidopteran green- and red-sensitive visual pigments form a monophyletic clade, which lends support to the hypothesis from comparative physiological studies that red-sensitive visual pigments in insects have paralogous origins. Polymorphic amino acid sites (180, 197, 277, 285, 308), which are essential for generating the spectral diversity among the vertebrate red- and green-sensitive pigments are notably invariant in the Papilio red- and green-sensitive pigments. Other major tuning sites must be sought to explain the spectral diversification among these and other insect visual pigments.

5' Untranslated Regions↗
PubMed2005

Physiological characterization of the compound eye in monarch butterflies with focus on the dorsal rim area.

The spectral, angular and polarization sensitivities of photoreceptors in the compound eye of the monarch butterfly (Danaus plexippus) are examined using electrophysiological methods. Intracellular recordings reveal a spectrally homogenous population of UV receptors with optical axes directed upwards and >or=10 degrees to the contralateral side. Based on optical considerations and on the opsin expression pattern (Sauman et al. 2005), we conclude that these UV receptors belong to the anatomically specialized dorsal rim area (DRA) of the eye. Photoreceptors in the main retina with optical axes <10 degrees contralateral or ipsilateral have maximal sensitivities in the UV (lambda(max)<or=340 nm), the blue (lambda(max)=435 nm) or in the long-wave range (green, lambda(max)=540 nm). The polarization sensitivity (PS) of the UV receptors in the DRA is much higher (PS=9.4) than in the UV cells (PS=2.9) or green cells (PS=2.8) of the main retina. The physiological properties of the photoreceptors in the DRA and in the main retina fit closely with the anatomy and the opsin expression patterns described in these eye regions. The data are discussed in the light of present knowledge about polarized skylight navigation in Lepidopterans.

Animals↗
PubMed2005

Visual pigment spectra of the comma butterfly, Polygonia c-album, derived from in vivo epi-illumination microspectrophotometry.

The visual pigments in the compound eye of the comma butterfly, Polygonia c-album, were investigated in a specially designed epi-illumination microspectrophotometer. Absorption changes due to photochemical conversions of the visual pigments, or due to light-independent visual pigment decay and regeneration, were studied by measuring the eye shine, i.e., the light reflected from the tapetum located in each ommatidium proximal to the visual pigment-bearing rhabdom. The obtained absorbance difference spectra demonstrated the dominant presence of a green visual pigment. The rhodopsin and its metarhodopsin have absorption peak wavelengths at 532 nm and 492 nm, respectively. The metarhodopsin is removed from the rhabdom with a time constant of 15 min and the rhodopsin is regenerated with a time constant of 59 min (room temperature). A UV rhodopsin with metarhodopsin absorbing maximally at 467 nm was revealed, and evidence for a blue rhodopsin was obtained indirectly.

Animals↗
PubMed2003

Evidence for Notch-mediated lateral inhibition in organizing butterfly wing scales.

Here I present gene expression data that implicate a Notch-mediated lateral inhibition process in the spatial organization of butterfly wing scales. During early pupal development the receptor molecule Notch is expressed in a grid-like pattern in the wing epithelium, resulting in parallel rows of uniformly spaced cells with low Notch expression. Previous work has shown that these low-Notch cells express a homolog of the achaete-scute proneural transcription factors and develop into scales. All of these observations are consistent with the Drosophila model of Notch-mediated bristle determination and support the hypothesis that bristles and scales share an underlying patterning mechanism.

Animals↗
PubMed2000

Butterfly wing pattern mutants: developmental heterochrony and co-ordinately regulated phenotypes.

Butterfly wings are colored late in development, when pigments are synthesized in specialized wing scale cells in a fixed developmental succession. In this succession, colored pigments are deposited first and the remaining areas are later melanized black or brown. Here we studied the developmental changes underlying two wing pattern mutants, firstly melanic mutants of the swallowtail Papilio glaucus, in which the yellow background is turned black, and secondly a Spotty mutant of the satyrid Bicyclus anynana, which carries two additional eyespots. Despite the very different pattern changes in these two mutants, they are both associated with changes in rates of scale development and correspondingly, the final color pattern. In the melanic swallowtail, background scales originally destined to become yellow (normally developing early and synthesizing papiliochrome) show delayed development, fail to make papiliochrome, and subsequently melanize at the same time as scales in the wild-type black pattern. In the B. anynana eyespot, scale maturation begins with the central white focus, then progresses to the surrounding gold ring and later finishes with melanization of the black center. Mutants showing additional eyespots display accelerated rates of scale development (corresponding to new eyespots) in wing cells not normally occupied by eyespots. Thus by either delaying or accelerating rates of scale development, the final color, or position, of a wing pattern element can be changed. We propose that this heterochrony of scale development is a basic mechanism of color pattern formation on which developmental mutants act to change lepidopteran color patterns.

Animals↗
PubMed2004

Adult stemmata of the butterfly Vanessa cardui express UV and green opsin mRNAs.

Adult stemmata are distinctive insect photoreceptors located on the posterior surfaces of the optic lobes. They originate as larval eyes that migrate inward during metamorphosis. We used a combination of light microscopy and in situ hybridization to examine their anatomical organization in the butterfly Vanessa cardui and to test for the presence of visual pigments, the light sensitive components of the visual transduction pathway. The bilateral cluster of six internal stemmata is located near the ventral edge of the lamina. They retain the dark screening pigment and overlying crystalline cones of the larval stemmata. We found two opsin mRNAs expressed in the stemmata that are also expressed, respectively, in UV-sensitive and green-sensitive photoreceptor cells in the compound eye. A third mRNA that is expressed in blue-sensitive photoreceptor cells of the compound eye was not expressed in the stemmata. Our results reinforce the idea that the adult stemmata are not merely developmental remnants of larval eyes, but remain functional, possibly as components of the circadian input channel.

Animals↗
PubMed2003

Winter warming facilitates range expansion: cold tolerance of the butterfly Atalopedes campestris.

Our ability to predict ecological and evolutionary responses to climate change requires an understanding of the mechanistic links between climate and range limits. The warming trend over the past half-century has generated numerous opportunities to develop much-needed case studies of these links. Species that are only limited by climatic factors are likely to shift range quickly during periods of warming. Such species directly impact recipient communities and indicate trends that will become more widespread. Because minimum temperature (T (min)) is rising at twice the rate of maximum temperature, species with this range-limiting factor may be especially responsive to global warming. In this study, I test the hypothesis that rising T (min) has directly affected the range of a skipper butterfly. Atalopedes campestris has moved northward rapidly this century, recently colonizing eastern Washington where January T (min) has risen 3 degrees C in 50 years. The results show that: 1. A. campestris' range lies completely within the -4 degrees C January average minimum isotherm, and that recently colonized areas were below this threshold earlier this century. 2. In acute cold stress experiments, -4 to -7 degrees C proved to be a critical thermal limit: median supercooling point was -6.3 degrees C, and minimum lethal temperature (LT50 with 12-h exposure) was -5.7 degrees C. 3. In chronic cold stress experiments, survivorship declined sharply in diurnally fluctuating thermal regimes typical of the current range edge. High mortality occurred under constant 0 degrees C conditions as well as in fluctuating regimes, implying that thermal insulation from snow would not protect A. campestris. 4. There was no evidence of evolution in cold tolerance at the range margin, despite strong selection. Thus, winter warming was apparently a prerequisite for the range expansion. Characteristics of this species that seem to be associated with its rapid response are that it is an opportunistic species, it is not habitat or dispersal limited, and it is constrained by T (min).

Acclimatization↗
PubMed2004

Field transplants reveal summer constraints on a butterfly range expansion.

The geographic ranges of most species are expected to shift to higher elevations and latitudes in response to global warming. But species react to specific environmental changes in individualistic ways, and we are far from a detailed understanding of range-shifts. Summer temperature often limits the ranges of insects and plants, so many range-shifts are expected to track summer warming. I explore this potential range-limiting factor in a case study of a northwardly expanding American butterfly, Atalopedes campestris (Lepidoptera, Hesperiidae). This species has recently colonized the Pacific Northwest, USA, where the mean annual temperature has risen 0.8-1.8 degrees C over the past 100 years. Using field transplant experiments across the current range edge, I measured development time, survivorship, fecundity and predation rates along a naturally occurring thermal gradient of 3 degrees C. Development time was significantly slower outside the current range in eastern Washington (WA), as expected because of cooler temperatures there. Slower development would reduce the number of generations possible per year outside the current range, dramatically lowering the probability that a population could survive there. Differences in survivorship, fecundity and predation rate across the range edge were not significant. The interaction between summer and winter temperature appears to be crucial in defining the current range limit. The estimated difference in temperature required to affect the number of generations is greater than the extent of summer warming observed over the past century, however, and thus historically winter temperature alone probably limited the range in southeastern WA. Nonetheless, extraordinarily warm summers may have improved colonization success, increasing the probability of a range expansion. These results suggest that extreme climatic events may influence rates of response to long-term climate change. They also demonstrate that range-limiting factors can change over time, and that the asymmetry in seasonal warming trends will have biological consequences.

Age Factors↗